Frogs of the Eleutherodactylus Rugulosus Group: A Cladistic Study of Allozyme, Morphological, and Karyological Data

The Eleutherodactylus rugulosus group of Savage (1975) contains 11 nominal species of frogs distributed throughout Mexico and Central America. Three subgroups are currently recognized from morphological data. The E. fleischmanni subgroup includes E. angelicus, E. escoces, E. fleischmanni, and E. punctariolus, whereas the E. rugulosus subgroup contains E. azueroensis, E. berkenbuschii, E. brocchi, E. rugulosus, E. taurus, and E. vocalis. These two subgroups form a mono-phyletic component related to the monotypic subgroup, E. merendonensis. A major purpose of the present study is to emphasize that phylogenetic relationships must be constructed with all available information subjected to cladistic methods. The tendency to emphasize one's own data to the complete or partial exclusion of others must be avoided in favor of synthetic approaches. Recently, the morphological data of Savage (1975) have been supplemented with allozyme and karyological characteristics. This recent availability of additional data provides the opportunity to evaluate the relationships of the E. rugulosus group based on a phylogenetic synthesis of multiple character sets. A general cladogram of the E. rugulosus group is constructed from a phylogenetic synthesis of allozyme, morphological, and karyological information. The allozyme, morphological, and karyological data are analyzed separately with the transformation series analysis procedure and the three sets of transformed results are combined into one character set. The combined data matrix is subjected to Wagner analysis and the most parsimonious tree is retained as the general cladogram. The hypotheses (phylogeny and classification) of Savage (1975) are evaluated and modified against the general cladogram. The phylogenetic relationships of the general cladogram do not support the hypotheses (phylogeny and classification) of Savage (1975). Eleutherodactylus azueroensis (the E. azueroensis subgroup) and E. vocalis (the E. vocalis subgroup) constitute separate lineages of a trichotomy originating from the root of the cladogram. All other species belong to the third lineage (the E. punctariolus subgroup) divided into two major components. These components include: (1) E. fleischmanni and the two sister species, E. angelicus and E. escoces (the E. fleischmanni infragroup); and (2) E. berkenbuschii, E. brocchi, E. merendonensis, E. punctariolus, E. rugulosus, and E. taurus (the E. punctariolus infragroup). Eleutherodactylus taurus and the two sister species, E. merendonensis and E. punctariolus, form a distinct cluster related to E. brocchi, E. berkenbuschii, and E. rugulosus in that order.